About the Authors

Michael M. Mendelson

Contributed equally to this work with: Michael M. Mendelson, Riccardo E. Marioni

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Boston University School of Medicine, Boston, Massachusetts, United States of America, Department of Cardiology, Boston Children’s Hospital, Boston, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Riccardo E. Marioni

Contributed equally to this work with: Michael M. Mendelson, Riccardo E. Marioni

Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, United Kingdom, Medical Genetics Section, Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, United Kingdom, Queensland Brain Institute, University of Queensland, Brisbane, Queensland, Australia

Roby Joehanes

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America, Hebrew SeniorLife, Harvard Medical School, Boston, Massachusetts, United States of America

Chunyu Liu

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America, Department of Biostatistics, Boston University, Boston, Massachusetts, United States of America

Åsa K. Hedman

Affiliation Molecular Epidemiology and Science for Life Laboratory, Department of Medical Sciences, Uppsala University, Uppsala, Sweden

Stella Aslibekyan

Affiliation Department of Epidemiology, School of Public Health, University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Ellen W. Demerath

Affiliation Division of Epidemiology and Community Health, School of Public Health, University of Minnesota, Minneapolis, Minnesota, United States of America

Weihua Guan

Affiliation Division of Biostatistics, School of Public Health, University of Minnesota, Minneapolis, Minnesota, United States of America

Degui Zhi

Affiliation Department of Biostatistics, School of Public Health, University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Chen Yao

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Tianxiao Huan

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Christine Willinger

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Brian Chen

Affiliation Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Paul Courchesne

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Michael Multhaup

Affiliation Center for Epigenetics, Johns Hopkins University School of Medicine, Baltimore, Maryland, United States of America

Marguerite R. Irvin

Affiliation Department of Epidemiology, School of Public Health, University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Ariella Cohain

Affiliation Icahn Institute for Genomics and Multiscale Biology and Department of Genetics and Genomic Sciences, Icahn School of Medicine at Mount Sinai, New York, New York, United States of America

Eric E. Schadt

Affiliation Icahn Institute for Genomics and Multiscale Biology and Department of Genetics and Genomic Sciences, Icahn School of Medicine at Mount Sinai, New York, New York, United States of America

Megan L. Grove

Affiliation Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, Texas, United States of America

Jan Bressler

Affiliation Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, Texas, United States of America

Kari North

Affiliation Department of Epidemiology, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

Johan Sundström

Affiliation Cardiovascular Epidemiology, Department of Medical Sciences, Uppsala University, Uppsala, Sweden

Stefan Gustafsson

Affiliation Molecular Epidemiology and Science for Life Laboratory, Department of Medical Sciences, Uppsala University, Uppsala, Sweden

Sonia Shah

Affiliations Queensland Brain Institute, University of Queensland, Brisbane, Queensland, Australia, Institute for Molecular Bioscience, University of Queensland, Brisbane, Queensland, Australia

Allan F. McRae

Affiliations Queensland Brain Institute, University of Queensland, Brisbane, Queensland, Australia, Institute for Molecular Bioscience, University of Queensland, Brisbane, Queensland, Australia

Sarah E. Harris

Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, United Kingdom, Medical Genetics Section, Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, United Kingdom

Jude Gibson

Affiliation Wellcome Trust Clinical Research Facility, Western General Hospital, University of Edinburgh, Edinburgh, United Kingdom

Paul Redmond

Affiliation Department of Psychology, University of Edinburgh, Edinburgh, United Kingdom

Janie Corley

Affiliation Department of Psychology, University of Edinburgh, Edinburgh, United Kingdom

Lee Murphy

Affiliation Wellcome Trust Clinical Research Facility, Western General Hospital, University of Edinburgh, Edinburgh, United Kingdom

John M. Starr

Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, United Kingdom, Alzheimer Scotland Dementia Research Centre, University of Edinburgh, Edinburgh, United Kingdom

Erica Kleinbrink

Affiliation Center for Molecular Medicine and Genetics and Department of Neurology, Wayne State University, Detroit, Michigan, United States of America

Leonard Lipovich

Affiliation Center for Molecular Medicine and Genetics and Department of Neurology, Wayne State University, Detroit, Michigan, United States of America

Peter M. Visscher

Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, United Kingdom, Queensland Brain Institute, University of Queensland, Brisbane, Queensland, Australia, Institute for Molecular Bioscience, University of Queensland, Brisbane, Queensland, Australia

Naomi R. Wray

Affiliations Queensland Brain Institute, University of Queensland, Brisbane, Queensland, Australia, Institute for Molecular Bioscience, University of Queensland, Brisbane, Queensland, Australia

Ronald M. Krauss

Affiliation Children’s Hospital Oakland Research Institute, Oakland, California, United States of America

Daniele Fallin

Affiliation Center for Epigenetics, Johns Hopkins University School of Medicine, Baltimore, Maryland, United States of America

Andrew Feinberg

Affiliation Center for Epigenetics, Johns Hopkins University School of Medicine, Baltimore, Maryland, United States of America

Devin M. Absher

Affiliation HudsonAlpha Institute for Biotechnology, Huntsville, Alabama, United States of America

Myriam Fornage

Affiliations Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, Texas, United States of America, Brown Foundation Institute of Molecular Medicine, University of Texas, Houston, Texas, United States of America

James S. Pankow

Affiliation Division of Epidemiology and Community Health, School of Public Health, University of Minnesota, Minneapolis, Minnesota, United States of America

Lars Lind

Affiliation Cardiovascular Epidemiology, Department of Medical Sciences, Uppsala University, Uppsala, Sweden

Caroline Fox

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Erik Ingelsson

Affiliations Molecular Epidemiology and Science for Life Laboratory, Department of Medical Sciences, Uppsala University, Uppsala, Sweden, Department of Medicine, Division of Cardiovascular Medicine, Stanford University School of Medicine, Stanford, California, United States of America

Donna K. Arnett

Affiliation College of Public Health, University of Kentucky, Lexington, Kentucky, United States of America

Eric Boerwinkle

Affiliations Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, Texas, United States of America, Human Genome Sequencing Center, Baylor College of Medicine, Houston, Texas, United States of America

Liming Liang

‡ These authors are joint senior authors on this work.

Affiliation Departments of Epidemiology and Biostatistics, School of Public Health, Harvard University, Boston, Massachusetts, United States of America

Daniel Levy

levyd@nih.gov

‡ These authors are joint senior authors on this work.

Affiliations Framingham Heart Study, Framingham, Massachusetts, United States of America, Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Ian J. Deary

‡ These authors are joint senior authors on this work.

Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, United Kingdom, Department of Psychology, University of Edinburgh, Edinburgh, United Kingdom

Competing Interests

JS is an advisory board member for Itrim. CF is now an employee of Merck, but was not an employee when the work was conducted. EI is a scientific advisor and consultant for Precision Wellness, Inc. and scientific advisor for Cellink for work unrelated to this paper. IJD has research grants from Age UK and the UK Medical Research Council.

Author Contributions

Conceptualization: MMM DL CF REM IJD PMV NRW EWD WG MLG JB JP EI MF EB DKA. Data curation: PR JC PC. Formal analysis: MMM REM RJ CL DZ BC SS AFM EWD WG MLG KN MF JP EK LLip AC EES ASK SG. Funding acquisition: DL IJD PMV NRW JMS. Investigation: MMM IJD JC JMS DKA LLin PC SEH JG LM WG MLG KN JP DMA MMu DF AF MRI JS JSP. Methodology: MMM REM EWD SA AKH LLin EJD RMK LLia DL IJD. Software: RJ LLia. Supervision: DL EI IJD EB MF DKA. Visualization: REM AKH CY TH CW. Writing – original draft: MMM. Writing – review & editing: REM RJ CL AKH SA EWD WG DZ CY TH CW BC PC MMu MRI AC EES MLG JB KN JS SG SS AFM SEH JG PR JC LM JMS EK LLip PMV NRW RMK DF AF DMA MF JSP LLin CF EI DKA EB LLia DL IJD.